Myrmecologische Nachrichten 8 59 - 68 Wien, September 2006
On the ants (Hymenoptera: Formicidae) of the Philippine Islands: I. The genus Pristomyrmex MAYR, 1866
Herbert ZETTEL
Abstract
The Philippine fauna of the ant genus Pristomyrmex MAYR, 1866 is analysed. Three species are described as new: Pristomyrmex distinguendus sp.n. from Luzon and Leyte, P. rugosus sp.n. from Leyte, and P. schoedli sp.n. from Leyte. One species, P. quadridens WANG, 2003, is recorded from the Philippines for the first time. Further unpublished records are presented for P. collinus WANG, 2003, P. longispinus WANG, 2003, P. picteti EMERY, 1893, and P. punctatus (F. SMITH, 1860). The genus contains 55 species world-wide. Seventeen species (= 30.9 %) are recorded from the Philip- pines, nine of which are endemic. An identification key to the Philippine species of Pristomyrmex is presented. Key words: Ants, Formicidae, Pristomyrmex, Philippines, key, new species, first record, fauna, endemism. Dr. Herbert Zettel, Natural History Museum, International Research Institute of Entomology, Burgring 7, A-1010 Vienna, Austria. E-mail: [email protected]
Introduction Pristomyrmex MAYR, 1866 is an easily recognised genus This paper adds three new species and one first re- of the Myrmecinini as defined by BOLTON (2003: 71). Re- cord from the Philippines, assigns old Philippine records cently, WANG (2003) revised the taxonomy of Pristomyr- to islands, provinces, and actual locality names, and ana- mex; this study also contains an overview on the history of lyses the Philippine Pristomyrmex fauna zoogeographical- the genus and a definition based on 30 morphological char- ly. The author also wants to encourage Filipino entomo- acteristics of the worker, and 27 of the male. logists to pay attention to these beautiful ants, which are in- Within the Philippine fauna, workers of Pristomyrmex teresting not only faunistically, but even more in ecolo- can be easily distinguished from other myrmicine ants by gical aspects: Most species seem to be indicators for un- the combination of the following characters: antenna 11- disturbed or weakly disturbed forests and thus might be segmented and without well defined club; antennal scrobes useful for habitat conservation measures. absent, or reduced and situated above the small eyes; an- terior clypeal margin denticulate or crenulate; mandible Material and methods distally broadened and with long axis rotated so that the All specimens are dry mounted on card squares or tri- masticatory margin is (almost) vertical; petiole peduncu- angles. Examination of specimens was carried out with a late (see BOLTON 1994). Most Philippine species of Pristo- LEICA Wild M10 binocular microscope; measurements myrmex have pronotal spines, a characteristic which is only were taken at magnifications of 80 × and 128 ×. Digital found in two other myrmicine genera occuring in the Phil- photographs were taken with a Leica DFC camera attached ippines, Acanthomyrmex EMERY, 1893 and Pheidole WEST- to a Leica MZ16 binocular microscope using Image Man- WOOD, 1839, both of which have 12-segmented antennae. ager IM50 and processed with Auto-Montage Pro and Only a small part of the material treated in the present Adobe Photoshop 7.0 programmes. study was identified by Wang (labelled in 1998) and includ- Terminology and method of description follow WANG ed in his excellent revision. These additional specimens were (2003), definitions of AL and TL2 follow WANG (2003: either collected later or had not been sorted at that time. fig. 2). The 55 species of Pristomyrmex are distributed in the Measurements and indices (see also WANG 2003: figs. tropics and subtropics from Africa to Japan and Australia 1-3): (see WANG 2003). The majority (36 species) are Oriental in HW Head width. Maximum width of head, in full-face the widest sense (incl. New Guinea), with one species ex- view in front of eyes (excluding eyes). tending its range to the southeastern Palaearctic Region. HL Head length, in full-face view, excluding mandibles Six of seven Australian species are endemic, as are all five measured from midpoint of a straight line across pre- African species, and three species on the island of Mauri- occipital margin either to frontal-most point of apex tius. As shown in this paper, the Philippines certainly is a of median tooth of anterior clypeal margin or, if centre of diversity for two reasons: First, species reached median tooth short or absent, to midpoint of line the Philippines from Sundaland and from the Papuan sub- connecting frontal-most apices of the two lateral region, both of which are very rich in species; secondly, teeth of anterior clypeal margin. there are several species which seem to be endemic on cer- CI Cephalic index. HW / HL * 100. tain islands. Seventeen species (= 30.9 % of world fauna; SL Scape length. Length of antennal scape, including including new species and records) inhabit the Philippines, lamella encircling base of scape but excluding basal nine of which are endemic (see Discussion and Tab. 1). condyle.
Stefan-Schödl-Gedenkband / Stefan Schödl Memorial Volume
SI Scape index. SL / HW * 100. formula, lamella of scape). Although the grouping is obvi- PW Pronotal width. Maximum width of pronotum in dor- ously useful for a phylogeny of the genus, it is less help- sal view. ful for this faunal survey. Instead, a key primarily based on AL Alitrunk length. Diagonal length of alitrunk in later- more easily recognizable characteristics is presented. al view, from frontal-most point of declivous area of pronotum to posterior-most point of apex of me- Pristomyrmex bicolor EMERY, 1900 tapleural lobe. Material examined: Luzon: Laguna Pr., Mt. Banahaw, EL Eye length. Maximum length of eye. above Kinabuhayan, trail to Crystalino, 24.XI.1995, leg. TL Total length = TL1 + TL2 + TL3. TL1: Line mea- J. Kodada & R. Rigová, det. Wang M., 1 ∑ (NHMW). sured from apex of closed mandibles to midpoint of Diagnosis of worker: TL 4.5 - 6.2 mm. Clypeus dor- a straight line across occipital margin, in full-face sally with strong median carina, ventrally variably modi- view. TL2: Straight line from declivous area of pro- fied, but always without distinct central tooth. Masticatory notum to point at which posterior margin of postpet- margin of mandible with diastema. Dorsum of head rugo- iole meets uppermost point of articulation of gaster. reticulate, of alitrunk smooth. Pronotal spines very long TL3: Line from anterior-uppermost point of arti- (PSL1 > 0.35, usually > 0.40), much longer than propo- culation to apex of gaster. deal spines. First gastral tergite without setae. PSL1 Pronotal spine length. Straight distance from ante- Previous records from the Philippines: Palawan rior base to apex of pronotal spine. (Binaluan) [types of P. taurus] (STITZ 1925); L u zo n : PSL2 Propodeal spine length. Straight distance from pos- Laguna (Mt. Banahaw) (WANG 2003). terior base to apex of propodeal spine. General distribution: Malay Peninsula, Java, Borneo, PPI Postpetiole index. PPW / PPL * 100. PPW: maxi- Philippines (WANG 2003). mum width of postpetiole in dorsal view. PPL: dor- Notes: The worker from Mt. Banahaw is the single so-median length of postpetiole. specimen of P. bicolor known from east of the Wallace' and All measurements are taken in millimetres. For para- Dickerson's Lines, which delimit the Pleistocenic Sunda types the minimum and maximum values are presented; n Shelf area to the east. It differs from other material exa- refers to the number of specimens measured. mined (including a syntype of P. bicolor) in a completely Locality data are arranged zoogeographically, within smooth pronotum and agrees in that characteristic with the Philippines the sequence follows the regions and sub- the type of P. taurus STITZ, 1925 from Palawan (synonymy regions listed by ONG & al. (2002) (see also Tab. 1). Data and discussion see WANG 2003: p. 425); also see discussion. are presented in the following order: (1) for Material exa- mined: "island, province, locality" for large islands con- Pristomyrmex brevispinosus EMERY, 1887 sisting of several administrative provinces, or "island, lo- No material from the Philippines examined. cality" for small island identical with or part of one pro- Diagnosis of worker: TL 3.0 - 4.3 mm. Clypeus dor- vince. (2) for Previous records: "island: province (locality / sally with median carina, ventrally with distinct central -ies)" for large islands consisting of several administrative tooth. Masticatory margin of mandible with diastema. Dor- provinces, or "island (locality)" for small island identical sum of head (at least behind eyes) and alitrunk rugoreti- with or part of one province. #-numbers are sample (loca- culate. Pronotal spines tooth-like (PSL1 0.06 - 0.12) sub- lity) numbers, but do not always refer to nest series. equal in length or slightly shorter than propodeal spines. Acronyms of repositories: First gastral tergite without setae. CZW Coll. H. & S.V. Zettel, Vienna, Austria Previous records from the Philippines: Mindanao: NHMW Natural History Museum, Vienna, Austria Bukidnon (Malaybalay) (WANG 2003). UPLB University of the Philippines, Los Baños, Philip- General distribution: Japan (YAMANE & TERAYAMA pines 1999); Taiwan, Thailand, Malay Peninsula, Sumatra, Bor- USC University of San Carlos (Entomological Collec- neo, Philippines, Sulawesi (WANG 2003). tion), Cebu City, Philippines Diagnoses of species are given for the purpose of dis- Pristomyrmex collinus WANG, 2003 tinguishing the Philippine species from each other. Speci- Material examined: Lu zo n : Laguna Pr., Los Baños, Mt. mens of some endemic species have not been available for Makiling, 13. - 18.XI.1992, leg. H. Zettel (# 1), det. Wang study, although the author asked for permission to study M., 1 ∑ (CZW); Laguna Pr., Los Baños, Mt. Makiling, Mud some paratype specimens. In such cases the diagnoses are Spring, 23. - 24.I.1999, leg. H. Zettel (# 167), 1 ∑ (CZW); based on the descriptions and diagnoses by WANG (2003), Laguna Pr., Mt. Banahaw, above Kinabuhayan, trail to and photographs of type specimens presented by ALPERT Crystalino, 24.XI.1995, leg. J. Kodada & R. Rigová, det. & al. (2006) were used to confirm some characteristics. Wang M., 3 ∑∑ (NHMW); Camarines Sur Pr., Lake Buhi area, nr. Itbog, Twin Falls, 22.III.1998, leg. H. Zettel Treatment of species (# 164), 3 ∑∑ (CZW); Tablas: San Agustin, Dubduban, The species are discussed in alphabetical order. According Busai Falls, 23. - 25.XI.1994, leg. H. Zettel (# 70), det. to WANG (2003), P. divisus, P. pulcher, and P. punctatus Wang M., 1 ∑ (CZW); Mindanao: Surigao del Norte Pr., belong to the P. punctatus species group; P. simplex to the 40 km S Surigao, Songkoy Spring, 8.II.2000, leg. S. Schödl P. levigatus species group; P. picteti to the P. umbripennis (# 11), 1 ∑ (NHMW). species group; and all remaining species (including the new Diagnosis of worker: TL 3.3 - 3.9 mm. HW 0.77 - species) to the P. quadridens species group. Some species 0.94. Clypeus ventrally with central tooth. Antennal scrobe group characteristics used by WANG (2003) are difficult to absent. Masticatory margin of mandible with diastema. recognize or to evaluate (e.g., dentation of mandibles, palp Dorsum of head and alitrunk smooth and punctate, with
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Figs. 1 - 2: Pristomyrmex distinguendus sp.n. (1) Head, full face view; (2) habitus, lateral view. sparse, relatively short setae. Pronotal spines tooth-like, sub- 4.III.1999, leg. F. Seyfert (# 19) (NHMW); paratypes: equal in length to propodeal spines. Anterior face of petio- Laguna Pr., Los Baños, Mt. Makiling, from UPLB to Mud lar node distinctly separated from dorsal surface of pe- Spring, 18.XI.1999, leg. H. Zettel (# 207), 1 ∑ (CZW); La- duncle. First gastral tergite without setae. guna Pr., Los Baños, Mt. Makiling, 500 - 1144 m, 14.XI. Previous records from the Philippines: Luzon: 1993, leg. H. Zettel (# 21a), P. collinus det. Wang M., 1 ∑ Laguna (Los Baños, Mt. Makiling, Mt. Banahaw), Rom- (CZW); Quezon Pr., Atimonan, Quezon NP, Old Zigzag blon Prov: T a b l a s (San Agustin), Panay (without fur- Road, 27. - 28.I.2002, leg. H. Zettel (# 300), 4 ∑∑ (CZW, ther information), Negros Oriental (Cuernos de Negros, UPLB); Leyte: Baybay, Leyte State University, c. 50 - Dumaguete [type locality]) (WANG 2003). – Luzon: Cam- 100 m, Calbiga-a River, 20. - 21.III.2005, leg. H. Zettel & arines Sur (Mt. Isarog Natural Park) (ALPERT & al. 2006). C. Pangantihon (# 422), 1 ∑ (CZW). General distribution: endemic to Philippines. Diagnosis of worker: Ventral surface of clypeus with strongly prominent tooth at centre. Pronotum with pair of Pristomyrmex costatus WANG, 2003 short teeth (PSL1 0.02 - 0.05), which shorter than propo- No material from the Philippines examined. deal spines (PSL2 0.05 - 0.09). Dorsum of head and ali- Diagnosis of worker: TL 4.1 - 4.6 mm. Clypeus dor- trunk smooth and polished, with many long hairs; dorsum sally with strong median carina, ventrally without central of head with scarce punctures (hair pits). Antennal scrobe tooth. Masticatory margin of mandible with diastema. Dor- present, its ventral margin defined by a ridge reaching at sum of head and alitrunk rugoreticulate. Pronotal spines least level of centre of eye; petiolar node with two or three moderately long (PSL1 0.19 - 0.27), distinctly longer than pairs of hairs. propodeal spines. First gastral tergite without setae. Measurements: Holotype worker: TL 3.32, HL 0.80, Previous records from the Philippines: Mindanao: HW 0.81, CI 101, SL 0.80, SI 100, EL 0.17, PW 0.58, AL Davao (Mt. McKinley) (WANG 2003). 0.87, PPW 0.26, PPL 0.27, PPI 96. – Paratype workers (n = General distribution: Malay Peninsula, Singapore, 7): TL 2.84 - 3.41, HL 0.68 - 0.81, HW 0.69 - 0.83, CI 100 Borneo, Philippines (WANG 2003). - 103, SL 0.70 - 0.85, SI 100 - 104, EL 0.15 - 0.19, PW 0.48 - 0.59, AL 0.75 - 0.90, PPW 0.21 - 0.28, PPL 0.23 - Pristomyrmex curvulus WANG, 2003 0.27, PPI 91 - 103. No material examined. Description of worker: Head (Fig. 1): Mandibles Diagnosis of worker: TL 4.6 - 5.3 mm. Clypeus dor- smooth and shiny. Masticatory margin of mandible with sally with median carina, ventrally with transverse ridge. four teeth: strongest apical + second strongest preapical + Masticatory margin of mandible with diastema. Dorsum of long diastema + two small basal teeth of similar size that head and alitrunk smooth and punctate. Pronotal spines are more or less fused. Basal margin of mandible almost very long (PSL1 > 0.35), longer than long propodeal straight, lacking distinct tooth. Clypeus with median lon- spines (PSL2 > 0.11). First gastral tergite without setae. gitudinal carina in caudal half (specimens from Luzon) or Previous records from the Philippines: Negros along entire length (specimen from Leyte). Anterior clypeus Oriental (Cuernos de Negros, Dumaguete [type locality]) margin with median tooth and one pair of lateral teeth. (WANG 2003). Ventral centre of clypeus with strongly prominent tooth. General distribution: endemic to Philippines. Palp formula 1, 3. Frontal carinae strong, extending caud- ad further than level of posterior eye margins. Antennal Pristomyrmex distinguendus sp.n. (Figs. 1, 2) scrobes present, laterally delimited by distinct ridge reach- Type material: holotype (worker): L u z o n : Camarines ing caudally at least to level of centre of eye. Frontal lobes Sur Pr., 20 km E Naga, E Carolina, slopes of Mt. Isarog, very weak; thus, antennal insertion almost entirely exposed.
61 Antennal scapes, when lying on dorsum of head, slightly Pristomyrmex hirsutus WANG, 2003 surpassing occipital margin of head. Eyes containing c. 8 - No material examined. 10 ommatidia in longest row. Profile shape of alitrunk and Diagnosis of worker: TL 5.8 mm. Clypeus dorsally pedicel segments as in Fig. 2. Pronotum with pair of short with reduced median carina, ventrally with small tooth. teeth, PSL1 0.02 - 0.05. Propodeum with pair of spines, Masticatory margin of mandible with long diastema. Dor- PSL2 0.05 - 0.09. Metapleural lobes subtriangular, acute. sum of head and alitrunk rugoreticulate. Pronotal spines Dorsum of alitrunk distinctly convex. Petiole in profile with short (PSL1 c. 0.10), propodeal spines tooth-like. Petiole fairly long peduncle, node with weakly developed angles, with high, dorsally rounded node. First gastral tergite with anterodorsal one on slightly higher level than posterodorsal numerous, evenly distributed, erect or suberect setae. one. Postpetiole in profile rounded dorsally, in dorsal view Previous records from the Philippines: Mindanao: broadening from front to back. Dorsum of head smooth Misamis Oriental ("Mt. Balatukan SW Gingoon" [type lo- and polished, with scattered hair pits which are small in cality]) (WANG 2003). specimens from Luzon and relatively large in specimen General distribution: endemic to Philippines. from Leyte; on scrobal areas, at least anteriorly, with some transverse rugae. Dorsum of alitrunk smooth and polished, Pristomyrmex longispinus WANG, 2003 with only very fine hair pits. Petiole, postpetiole, and gas- ter smooth and shiny. Dorsal surfaces of the head and ali- Material examined: Negros: Negros Oriental Pr., Cuer- trunk with numerous long erect or suberect hairs. Dorsum nos de Negros, Valencia, Apolong, Casaroro Falls, 25. - of petiolar node with 2 - 3 pairs of hairs; dorsum of post- 26.X.2004, leg. C. Pangantihon (# P400), 5 ∑∑ (CZW); petiole with 1 - 2 pairs. First gastral tergite lacking erect or ibid., 25. - 26.X.2004, leg. H. Zettel (# 400), 2 ∑∑ (NHMW); suberect hairs. A few pairs of forward projecting hairs pre- ibid., 9. - 13.III. 2005, leg. H. Zettel (# 420), 2 ∑∑ (CZW); sent near anterior clypeal margin. Scapes and tibiae with ibid., 26.III.2006, leg. H. Zettel & C. Pangantihon (# 451), numerous erect to suberect hairs. Colour reddish-brown. 6 ∑∑ (CZW). Comparative notes: This species is similar to P. colli- Diagnosis of worker: TL 4.5 - 5.7 mm. Clypeus dor- nus, but differs in a good number of characteristics: The sally without median carina, ventrally with transverse ruga. dorsal surface of the head and the alitrunk has many long Masticatory margin of mandible with diastema. Dorsum of setae. The pronotal spines are distinctly shorter than the pro- head and alitrunk smooth, on head sparsely punctate. Pro- podeal spines. The antennal scrobes are present and late- notal spines very long (PSL1 0.40 - 0.50) about twice as rally delimited by a ridge reaching caudally at least to level long as propodeal spines (PSL2 0.18 - 0.26). Petiolar node of centre of eye. The clypeus has three teeth, with the me- not separable from peduncle. First gastral tergite without dian tooth longest (observed also in one specimen of P. setae. collinus, in which dentation of clypeus is more variable Previous records from the Philippines: Negros than in P. distinguendus sp.n.). Oriental (Cuernos de Negros, Dumaguete [type locality]) General distribution: endemic to Philippines. (WANG 2003). General distribution: endemic to Philippines. Pristomyrmex divisus WANG, 2003
No material examined. Pristomyrmex picteti EMERY, 1893 Diagnosis of worker: TL 3.0 - 3.4 mm. Eye of mode- rate size, 8 - 10 ommatidia in longest row. Frontal carina Material examined: Luzon: Laguna Pr., Los Baños, Mt. short, not extending to level of posterior margin of eye. Makiling, 150 - 600 m, 13. - 14.XI.1993, leg. H. Zettel (# Lateral portions of clypeus not reduced. Masticatory mar- 21), det. Wang M., 1 dealate gyne (CZW). Leyte: Leyte gin of mandible with diastema. Dorsum of head foveolate, Pr., N Tacloban, Babatngon, Busay Falls, 28.I.2000, leg. H. of alitrunk smooth and with some rugae. Pronotal spines Zettel (# 220), 1 alate gyne (CZW); Leyte Pr., Baybay, Mt. absent, propodeal spines long. Petiole with long peduncle Pangasugan, Calbiga-a River, 50 - 200 m, 12.II.2000, leg. not clearly separated from short and high node. H. Zettel (# 236), 1 ∑ (CZW). Previous records from the Philippines: Negros Diagnosis of worker: TL 4.1 - 5.9 mm. Eyes very Oriental (Dumagete [type locality]) (WANG 2003). small, with 3 - 4 ommatidia in longest row. Clypeus dor- General distribution: endemic to Philippines. sally with median carina, ventrally with transverse ruga. Masticatory margin of mandible without diastema. Dorsum Pristomyrmex flatus WANG, 2003 of head and alitrunk foveolate. Pronotal spines absent, pro- No material examined. podeal spines relatively short. Node of petiole longer than Diagnosis of worker: TL 3.7 - 4.2 mm. HW 0.98 - high, well separated from peduncle. 1.04. Clypeus dorsally with indistinct median carina (only Previous records from the Philippines: Palawan posteriorly indicated), ventrally with weak medial tuber- (Binaluan) (STITZ 1925); – Luzo n : Ilocos Norte (Banqui cle. Masticatory margin of mandible with diastema. Dorsum = "Bauqui"), Rizal (Montalban), Laguna (Los Baños, Mt. of head and alitrunk smooth, on head sparsely punctate. Makiling, Mt. Banahaw); Palawan (Binaluan, Puerto Prin- Pronotal spines tooth-like, propodeal spines slightly lon- cesa); Negros Oriental (Cuernos de Negros, Dumaguete); ger. Anterior face of petiolar node not separated from dor- M i n d a n a o : Davao (Mt. Apo) (WANG 2003); – Lu zon : sal surface of peduncle. First gastral tergite without setae. Camarines Sur (Mt. Isarog Natural Park) (ALPERT & al. Previous records from the Philippines: Luzon: Ilocos 2006). Norte (Banqui = "Bauqui" [type locality]) (WANG 2003). General distribution: Malay Peninsula, Singapore, General distribution: endemic to Philippines. Phili ppines, New Guinea (WANG 2003).
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Figs. 3 - 4: Pristomyrmex rugosus sp.n. (3) Head, full face view; (4) habitus, lateral view.
Pristomyrmex pulcher WANG, 2003 Pristomyrmex quadridens EMERY, 1897 No material examined. Material examined: L e y t e : Baybay, Leyte State Univer- Diagnosis of worker: TL 2.7 - 3.1 mm. Lateral portions sity, c. 50 m, Lago-Lago River, 19.III.2005, leg. H. Zettel of clypeus not reduced. Masticatory margin of mandible & C. Pangantihon (# 421), 44 ∑∑ (CZW, NHMW, USC). with diastema, basal margin with broad tooth. Dorsum of First record for the Philippines! head and alitrunk rugoreticulate. Pronotal spines absent, Diagnosis of worker: TL 3.3 - 4.1 mm. Clypeus ven- propodeal spines long. Petiole almost triangular in lateral trally with median tooth. Masticatory margin of mandible view, peduncle not clearly separated from node. Tibiae yel- with diastema. Dorsum of head and alitrunk smooth and low. sparsely foveolate; foveae on head large. Pronotal spines Previous records from the Philippines: Camiguin tooth-like about as long as propodeal spine. First gastral (without further information) (ALPERT & al. 2006). tergite without setae. General distribution: Malay Peninsula (WANG 2003), General distribution: Previously known from New Philippines (ALPERT & al. 2006). Guinea and some nearby islands (Ceram, Ambon, etc.); WANG (2003: fig. 162) postulated an allopatric distribu- Pristomyrmex punctatus (F. SMITH, 1860) tion of P. quadridens and the closely related species P. Material examined: Marinduque: 1 km N Sihi, Mali- brevispinosus, which is also recorded from the Philippines. nao Spring, 16.II.1998, leg. H. Zettel (# 139), 1 ∑ (NHMW). As shown in this study, the two species have an overlap- Camotes: Pacijan Island, San Francisco, Northern Pob- ping distribution in the region "Greater Mindanao". lacion, Lake Danao, 27.II.2001, leg. H. Zettel (# 281), 1 ∑ (CZW); L e y t e : Leyte Prov., Hilusig, 14.II.2000, leg. H. Pristomyrmex rugosus sp.n. (Figs. 3, 4) Zettel (# 238), 1 ∑ (CZW). Bohol: Loboc, bank of Loboc Type material: holotype (worker): Leyte: Leyte Pr., Bay- River at Loboc Falls, 27.XI.2005, leg. C. Pangantihon (# bay, Mt. Pangasugan, Calbiga-a River, 50 - 200 m, 12.II. P424a), 23 ∑∑ (CZW, NHMW); same locality and date, 2000, leg. H. Zettel (# 236) (NHMW); paratypes: same data, leg. H. Zettel (# 424a), 4 ∑∑ (NHMW). Mindanao: Suri- 5 ∑∑ (CZW); same locality, c. 50 - 100 m, Calbiga-a River, gao del Norte, 40 km S Surigao, Songkoy Spring, 8.II.2000, 20. - 21.III.2005, leg. H. Zettel & C. Pangantihon (# 422), leg. H. Zettel (# 231), 16 ∑∑ (CZW); same locality and 2 ∑∑ (CZW, USC). date, leg. S. Schödl (# 11), 6 ∑∑ (NHMW); Surigao del Diagnosis of worker: Ventral surface of clypeus with Norte, SW Bacuag, Payapag, "Little Baguio" Waterfalls, prominent tooth at centre. Pronotum with pair of long spines 6.II.2000, leg. H. Zettel (# 228), 1 ∑ (CZW). (PSL1 0.15 - 0.17), which are distinctly longer than short Diagnosis of worker: TL 2.6 - 3.3 mm. Lateral por- propodeal teeth (PSL2 0.04 - 0.05). Dorsum of head and tions of clypeus reduced. Masticatory margin of mandible alitrunk with foveolate-reticulate sculpture or rugoreticu- with diastema, basal margin with or without inconspicu- letum, with long, relatively dense pilosity; petiolar node ous tooth. Dorsum of head and alitrunk rugoreticulate. Pro- and postpetiolar nodes smooth, each with several pairs of notal spines absent, propodeal spines long. Petiole with pe- hairs on dorsum. First gastral tergite without erect or sub- duncle not clearly separated from node. Tibiae brown or erect hairs. reddish. Measurements: Holotype worker: TL 4.26, HL 1.05, Previous records from the Philippines: Mindoro HW 1.02, CI 98, SL 1.09, SI 106, EL 0.22, PW 0.72, AL Occidental (San José), S a m a r (without further informa- 1.14, PPW 0.31, PPL 0.34, PPI 89. – Paratype workers (n = tion), Mindanao: Agusan (Talacogon) (WANG 2003). 7): TL 4.01 - 4.26, HL 0.94 - 1.05, HW 0.94 - 1.02, CI 97 - General distribution: southern and central China, Ja- 101, SL 1.01 - 1.06, SI 102 - 107, EL 0.19 - 0.21, PW 0.63 pan, Taiwan, Thailand, Vietnam, Singapore, Borneo, Phil- - 0.70, AL 1.00 - 1.12, PPW 0.27 - 0.30, PPL 0.30 - 0.32, ippines, New Guinea (WANG 2003). PPI 90 - 97.
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Description of worker: Head (Fig. 3): Mandibles more or less striate. Masticatory margin of mandible with four teeth: strongest apical + second strongest preapical + long diastema + two small basal teeth of similar size. Basal margin of mandible almost straight, lacking distinct tooth. Clypeus with strong median longitudinal carina and one pair of more or less developed lateral carinae. Anterior cly- peus margin with median tooth and three pairs of lateral teeth. Ventral centre of clypeus with strongly prominent tooth. Palp formula 1, 3. Frontal carinae strong, extending caudad further than level of posterior eye margins. Anten- nal scrobes present, laterally delimited by distinct ridge reaching caudally at least to level of centre of eye. Both frontal carinae and lateral ridge confluent with strong rugo- reticulum of head. Frontal lobes absent, antennal insertion entirely exposed. Antennal scapes, when lying on dorsum of head, surpassing occipital margin of head. Eyes con- taining 10 - 12 ommatidia in longest row. Profile shape of alitrunk and pedicel segments as in Fig. 4. Pronotum with pair of relatively long spines, PSL1 0.15 - 0.17. Propodeum with pair of teeth, PSL2 0.04 - 0.05. Metapleural lobes sub- triangular, apex weakly to moderately rounded. Dorsum of alitrunk anteriorly convex, posteriorly straight. Petiole in profile with fairly long peduncle, node with weakly devel- oped, high anteriodorsal angle, posteriodorsally rounded. Postpetiole in profile rounded dorsally, in dorsal view broadening from front to back. Dorsum of head, except for transversely rugous scrobal areas, and alitrunk with coarse rugoreticulum. Sides of pronotum similarly reticulate, but rugae less high. Petiole, postpetiole, and gaster smooth and shiny. Dorsal surfaces of head, alitrunk, petiole, and post- petiole with numerous long erect or suberect hairs. First gastral tergite lacking erect or suberect hairs. A few pairs of forward projecting hairs present near anterior clypeal margin. Scapes and tibiae with numerous erect to suberect Fig. 5: Pristomyrmex schoedli sp.n., habitus, dorsal view hairs. Colour dark reddish-brown. (Matthias Buch pinxit). Comparative notes: This species is most similar to P. sulcatus, but differs in having relatively long pronotal paratypes: same data, 119 ∑∑ (CZW, UPLB, USC); Leyte spines, dorsally more rounded node of petiole, and rela- Pr., Baybay, VISCA, 50 m, above Forestry Department, tively large eyes with more ommatidia (10 - 12 vs. 6 - 7 in stream, 11.II.2000, leg. H. Zettel (# 235), 2 ∑∑ (CZW); P. sulcatus fide WANG 2003). From typical P. sulcatus, it same locality and date, leg. S. Schödl (# 14), 2 ∑∑ (NHMW); can be easily distinguished by the numerous setae on the Leyte Pr., Baybay, VISCA, 50m, 31.I.2000\ leg. H. Zettel petiole and postpetiole and by the more reticulate dorsum (# 222), 1 ∑ (CZW); Baybay, Leyte State University, c. of the pro-mesonotum (with coarser longitudinal ridges in 50 m, Lago-Lago River, 19.III.2005, leg. H. Zettel & C. Pa- syntype of P. sulcatus). However, judging from the spe- ngantihon (# 421), 5 ∑∑ (CZW). cimens deposited in NHMW and identified by M. Wang, P. Diagnosis of worker: Ventral surface of clypeus with sulcatus as redescribed and interpreted by WANG (2003) strongly prominent tooth at centre. Clypeus without medi- is a variable species and may include further unrecognized an carina, anterior margin usually with seven (rarely five) taxa. From three other species, which are recorded from small denticles. Pronotum with pair of very long spines the Philippines, the new species differs as follows: from (PSL1 0.38 - 0.44), which are much longer than propodeal P. bicolor in the much shorter pronotal spines, in smaller spines (PSL2 0.09 - 0.13). Dorsum of head and alitrunk size, and the presence of a central tooth on the ventral sur- smooth and polished; dorsum of head with scarce punc- face of the clypeus; from P. brevispinosus in the abso- tures (hair pits). Frontal carinae posteriorly faded; antennal lutely and relatively longer pronotal spines; and from P. scrobes indistinct. Anterior face of petiolar node distinct costatus in the presence of a central tooth on the ventral sur- from dorsal surface of peduncle. Petiolar node with two or face of the clypeus, slightly shorter pronotal spines, and three pairs of hairs. more coarse rugoreticulum on dorsum of alitrunk. Measurements: Holotype worker: TL 4.44, HL 1.04, General distribution: endemic to Philippines. HW 1.15, CI 110, SL 1.17, SI 102, EL 0.22, PW 0.70, AL 1.05, PPW 0.30, PPL 0.32, PPI 94. – Paratype workers (n = Pristomyrmex schoedli sp.n. (Figs. 5 - 7) 10): TL 4.19 - 3.41, HL 0.99 - 1.10, HW 1.12 - 1.23, CI Type material: holotype (worker): L e y t e : Baybay, Leyte 113 - 121, SL 1.19 - 1.28, SI 103 - 107, EL 0.21 - 0.26, State University, c. 50 - 100 m, Calbiga-a River, 20. - 21.III. PW 0.72 - 0.75, AL 1.04 - 1.16, PPW 0.29 - 0.32, PPL 2005, leg. H. Zettel & C. Pangantihon (# 422) (NHMW); 0.30 - 0.37, PPI 104 - 118.
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Figs. 6 - 7: Pristomyrmex schoedli sp.n. (6) Head, full face view; (7) habitus, lateral view.
Description of worker: Head (Fig. 6): Mandibles Comparative notes: This species is similar to P. cur- smooth and shiny, with or without a few longitudinal rugae. vulus, but differs in the absence of a median carina on the Masticatory margin of mandible with four teeth: strong- clypeus and in the reduced frontal carina. From P. longi- est apical + second strongest preapical + long diastema + spinus this species differs in the shape of the petiole, in two small basal teeth of similar size. Basal margin of man- the relatively short propodeal spines and in shorter pilosity. dible almost straight, lacking tooth. Clypeus short, without The cephalic index of P. schoedli sp.n. (113 - 221) is median longitudinal carina. Anterior clypeus margin with slightly larger than in P. curvulus and P. longispinus (97 - short median tooth and three (rarely two) pairs of short 105 and 103 - 109; data from WANG 2003). lateral denticles. Ventral centre of clypeus with prominent General distribution: endemic to Philippines. tooth. Palp formula 1, 3. Frontal carinae anteriorly strongly developed, posteriorly indistinct and extending caudad at Pristomyrmex simplex WANG, 2003 most to level of posterior eye margins, but usually shor- Material examined: L u z o n : Laguna Pr., Mt. Banahaw, ter. Antennal scrobes indistinct, laterally not delimited. above Kinabuhayan, trail to Crystalino, 24.XI.1995, leg. Frontal lobes absent; antennal insertion entirely exposed. J. Kodada & R. Rigová, det. Wang M., 1 ∑ (NHMW). Frons anteriorly with very fine median impression, variab- Diagnosis of worker: TL 2.4 - 2.7 mm in types from ly reduced to short longitudinal groove. Antennal scapes, New Guinea, 3.0 mm in Philippine specimen. Eyes small. when lying on dorsum of head, slightly surpassing occi- Anterior clypeus margin with medial denticle and pair of pital margin of head. Eyes containing c. 10 - 12 ommati- strong lateral teeth. Masticatory margin of mandible with- dia in longest row. Profile shape of alitrunk and pedicel out diastema. Dorsum of head and alitrunk foveolate. Pro- segments as in Fig. 7. Pronotum with pair of long spines, notal spines absent, propodeal spines tooth-like. Petiolar PSL1 0.38 - 0.44. Propodeum with pair of short spines, node high, well separated from short peduncle. PSL2 0.09 - 0.13, distinctly shorter than distance between Remarks: The single worker from the Philippines is apices. Dorsum of alitrunk somewhat flattened, behind pro- considerably larger than the type specimens from New notal spines almost straight in lateral view. Metapleural Guinea and was not included in the type material by WANG lobes almost rectangular, with acute or weakly rounded (2003). apex. Petiole in profile with long peduncle; peduncle dis- Previous records from the Philippines: Luzon: tinct from anterior surface of node; node with weakly de- Laguna (Mt. Banahaw) (WANG 2003). veloped angles, anterodorsal one on slightly higher level General distribution: New Guinea, Philippines (WANG than posterodorsal one. Postpetiole in profile rounded dor- 2003). sally, in dorsal view slightly broadening from front to back. Dorsum of head smooth and polished, with some very fine hair pits; scrobal areas anteriorly with few transverse wrin- Identification key to the Philippine species of Pristomyrmex (workers only) kles. Dorsum of alitrunk smooth and polished, with only a few, very fine hair pits. Petiole, postpetiole, and gaster 1 Pronotal spines not at all developed...... 2 smooth and shiny. Dorsal surfaces of head and alitrunk with – Pronotal spines developed, short and triangular numerous long erect or suberect hairs. Dorsum of petio- or long (Figs. 2, 4, 5, 7)...... 6 lar node with 2 - 3 pairs of hairs; dorsum of postpetiole with several pairs. First gastral tergite lacking erect or sub- 2 Dorsal surface of head and alitrunk rugoreticu- erect hairs. A few pairs of forward projecting hairs pre- late...... 3 sent near anterior clypeal margin. Scapes and tibiae with – Dorsal surface of head and alitrunk punctate or numerous erect to suberect hairs. Colour reddish-brown. foveolate...... 4
65 3 Femora and tibiae reddish or brown; basal mar- least to level of centre of eye (Fig. 1)...... gin of mandible with or without inconspicuous ...... P. distinguendus sp.n. tooth...... P. punctatus – Pronotal spines subequal in length to propo- – Femora and tibiae yellow; basal margin of man- deal spines; antennal scrobes absent, laterally dible with a broad tooth...... P. pulcher not delimited by ridge...... P. collinus 4 Masticatory margin of mandible without dia- 15 Clypeus with median carina...... P. curvulus stema; petiolar node in profile longer than high – Clypeus without median carina (Fig. 6)...... 16 and distinct from peduncle...... P. picteti 16 Petiolar node in profile lacking distinct anterior – Masticatory margin of mandible with diastema; surface separated from dorsal surface of pe- petiolar node in profile higher than long or not duncle; propodeal spines relatively long (PSL2 clearly separated from peduncle...... 5 0.18 - 0.26)...... P. longispinus 5 Frontal carina short, not extending to level of – Petiolar node in profile with distinct anterior posterior margin of eye; masticatory margin of surface separated from dorsal surface of pedun- mandible with diastema...... P. divisus cle (Fig. 7); propodeal spines relatively short – Frontal carina long, extending to level of pos- (PSL2 0.09 - 0.13)...... P. schoedli sp.n. terior margin of eye; masticatory margin of mandible without diastema...... P. simplex Discussion 6 Dorsal surface of head (and usually also alitrunk) Taxonomy and morphology rugoreticulate (Figs. 3, 4)...... 7 Since the revision by WANG (2003) the taxonomy of Pris- – Dorsal surface of head and alitrunk smooth or tomyrmex is well known. However, a few of the widely punctate (Figs. 1, 2, 5 - 7)...... 11 distributed "species" may in fact be assemblages of highly similar species. One of them is P. sulcatus, a relative of 7 First gastral tergite with numerous, evenly dis- P. rugosus sp.n. (see comparative notes of that species). tributed, erect or suberect setae...... P. hirsutus The synonymy of P. taurus and P. bicolor (WANG 2003) – First gastral tergite without erect or suberect needs be re-examined, as soon as more material from the setae (Fig. 4)...... 8 Philippines becomes available. Pristomyrmex collinus sen- 8 Ventral surface of clypeus without central tooth...... 9 su WANG (2003) also consists of at least two species, be- cause one paratype of P. distinguendus sp.n. has been – Ventral surface of clypeus with a central tooth. .. 10 identified as P. collinus by M. Wang. As shown in the com- 9 Pronotal spines very long (at least 0.36 mm), parative notes of P. distinguendus sp.n., these two species usually longer than distance between their bases. are easily separable by several characters. The single para- ...... P. bicolor type of P. distinguendus sp.n. from Leyte differs slightly – Pronotal spines short (shorter than 0.30 mm), from material originating in Luzon, but is presently con- much shorter than distance between their bases. sidered conspecific. The single specimen of P. simplex ...... P. costatus known from the Philippines is much larger than the types from New Guinea and differs in a number of additional 10 Pronotal spines very short (0.06 - 0.12 mm), at characteristics (see WANG 2003); more material is required most as long as propodeal spines. . P. brevispinosus to confirm or reject conspecificity. – Pronotal spines relatively long (0.15 - 0.17 mm), In general, intraspecific variability appears to be very distinctly longer than propodeal spines (Fig. 4). low in species of Pristomyrmex, which makes identifica- ...... P. rugosus sp.n. tion relatively simple compared to many other myrmicine genera. 11 Pronotal spines triangular, short (compare Fig. 2)...... 12 Ecology – Pronotal spines long (compare Fig. 7)...... 15 Most species of Pristomyrmex dwell in the rainforest, for- 12 Dorsum of head with large foveolate punctures. aging as predators or scavengers (WANG 2003). Of the spe- ...... P. quadridens cies collected by the author, only P. punctatus has been found in obviously disturbed habitats; this species is the – Dorsum of head with relatively fine punctures most widespread in the genus (see WANG 2003). All other (Fig. 1)...... 13 Philippine species seem to be confined to humid forest hab- 13 Petiolar node in profile without distinct anteri- itats. Typical collection sites are wet, mossy rock faces or or face separated from upper surface of pedun- fallen tree trunks. Rarely, specimens have been observed cle; head width 0.98 - 1.04 mm...... P. flatus foraging on leaves (a few specimens of P. longispinus and P. schoedli). A nest of P. quadridens has been discovered – Petiolar node in profile with anterior face dis- in a piece of rotten wood (dimensions about 15 × 20 × tinctly separated from upper surface of pedunc- 50 cm) laying on the moist soil and rocks on a river bed. le; head width 0.77 - 0.94 mm...... 14 Usually specimens of Pristomyrmex are only collected in 14 Pronotal spines distinctly shorter than propode- small numbers, but approximately 100 P. schoedli workers al spines (Fig. 2); antennal scrobes present, lat- were collected within 10 minutes from a fallen log at Bay- erally delimited by ridge reaching caudally at bay (site # 422) and numerous specimens remained un-
66 Tab. 1: Checklist of Philippine species of Pristomyrmex, with distribution in biogeographical regions (following ONG & al. 2002). Abbreviations: A: Batanes; B: Babuyanes; C: Greater Luzon; D: Lubang; E: Greater Mindoro; F: Greater Pala- wan; G: Burias; H: Sibuyan; I: Romblon-Tablas; J: Greater Negros-Panay; K: Greater Mindanao; L: Camotes; M: Siquijor; N: Camiguin; O: Greater Sulu; P: Sibutu; + = species occurs also outside the Philippines; e = species endemic to Philippines; ? record may belong to an undescribed species.
Species A B C D E F G H I J K L M N O P +/e
P. bicolor EMERY, 1900 C F +
P. brevispinosus EMERY, 1887 K +
P. collinus WANG, 2003 C I J K e
P. costatus WANG, 2003 K +
P. curvulus WANG, 2003 J e
P. distinguendus sp.n. C K e
P. divisus WANG, 2003 J e
P. flatus WANG, 2003 C e
P. hirsutus WANG, 2003 K e
P. longispinus WANG, 2003 J e
P. picteti EMERY, 1893 C F J K +
P. pulcher WANG, 2003 N +
P. punctatus (F. SMITH, 1860) C E K L +
P. quadridens EMERY, 1897 K +
P. rugosus sp.n. K e
P. schoedli sp.n. K e
P. simplex WANG, 2003 ?C? +
number of species / region 0 0 7 0 1 2 0 0 1 5 10 1 0 1 0 0
collected, which suggests that nest sizes in Pristomyrmex fined by ONG & al. 2002). In this context it is interesting can occasionally be high. Low worker numbers, large dis- to note that most of the endemic species (except P. col- tance between specimens running on trails, and the rela- linus and P. distinguendus) are so far only recorded from tively slow motion of undisturbed specimens make Pristo- one region, but four of the eight non-endemic species are myrmex rather discreet animals. Nevertheless, Pristomyrmex recorded from two to four regions in the Philippines (see seem to be rare organisms. Deforestation on many Philip- Tab. 1). Whether this fact is influenced by rarity of some pine islands may already have reduced the diversity of species or reflects real regional endemism should be inves- those species, which are restricted to moist forests. tigated in future. Most species records are from Greater Mindanao (10 species), Greater Luzon (7 species), and Species diversity and regional endemism in the Greater Negros-Panay (5 species) (Tab. 1), probably be- Philippines cause of the more intensive collecting activities and more At present, 55 species of Pristomyrmex are described (WANG suitable habitats on the larger islands. 2003, and this paper). Seventeen species are recorded from the Philippines (= 30.9 % of world fauna). Of these seven- Acknowledgements teen species, nine species seem to be endemic to the Phil- My first thanks are addressed to the late Stefan Schödl for ippines (= 16.4 % or one sixth of world fauna). The ques- many enjoyable days together in the museum and in the tion, whether species are regionally endemic or not, needs field, when we could discuss ant taxonomy and diversity to be answered with caution due to the relatively small num- among many other topics. Stefan's ideas as well as his col- ber of collections. Table 1 presents records of the species lections in the Philippines contribute much to the present in the biogeographical regions of the Philippines (as de- study. Additional specimens used for this study have been
67 provided by J. Kodada and R. Rigová (Bratislava), C. Pa- References ngantihon (USC, Cebu City), and F. Seyfert (Vienna). My ALPERT, G.D., COVER, S., GENERAL, D.M. & SAMARITA, V. 2006: entomological research in the Philippines is supported by Ants of the Philippines. –
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